Table Of Content2004. The Journal of Arachnology 32:109-186
THE SYSTEMATICS OF SOUTHERN AFRICAN PARABUTHUS
POCOCK (SCORPIONES, BUTHIDAE): REVISIONS TO THE
TAXONOMY AND KEY TO THE SPECIES
Lorenzo Prendini: Division of Invertebrate Zoology, American Museum of Natural
History, Central Park West at 79th Street, New York, NY 10024-5192, USA. E-mail:
@
lorenzo amnh.org
ABSTRACT. The southern African species of Parabuthus Pocock 1890 are reviewed. Twenty valid
species are recognized and an illustrated key is provided for their identification. The diagnosis of each
species is revised, its known distributional range summarized, and notes on its ecology and conservation
provided. Three species areredeseribed andtheirdistributions mapped: P. calvusPurcell 1898;P. capensis
(Ehrenberg 1831); P. planicauda (Pocock 1896). Six new synonyms are proposed: Buthus brevimanus
van (3 segnis Thorell 1876 = P. granulatus (Ehrenberg 1831); Buthus villosus van p dilutus Thorell 1876
= P. raudus (Simon 1888); P granulatus strenuus Hewitt 1918 = P granulatus (Ehrenberg 1831); P.
capensisfrenchi Purcell 1901.= P. planicauda (Pocock 1889); Scorp.io teter Muller 1828 = P. trans-
vaalicus Purcell 1899; P. brachystylus Lawrence 1928 — P. villosus (Peters 1862). Two species, synon-
ymized by previous authors, but subsequently resurrected, are returned to synonymy: P. neglectus Purcell
1899 = P. capensis (Ehrenberg 1831), first synonymized by Kraepelin (1908); P. flavidus Pocock 1899
= P. mossambicensis (Peters 1861), first synonymized by Kraepelin (1914).
Keywords: Scorpiones, Buthidae, Parabuthus taxonomy, identification, distribution, ecology
,
mm
Parabuthus Pocock 1890 is an exclusively 600 of annual rainfall (Newlands 1978a,
Old World genus of scorpions, one of 82 gen- b).
era in the diverse, cosmopolitan family Buth- The dependence of Parabuthus species on
idae (Fet & Lowe 2000; Kovafik 2001, 2002, aridity has some fortunate implications for the
2003a). The genus includes the world's largest geographical distribution of scorpionism in
buthid scorpions, e.g., P. granulatus (Ehren- southern Africa, as Parabuthus includes some
berg 1831), P. schlechteri Purcell 1899, P. of the world’s most dangerously venomous
transvaalicus Purcell 1899 and P. villosus scorpions. Envenomation by these scorpions
(Peters 1862), which reach lengths of up to is a significant cause of morbidity and, in
mm
140 and masses of up to 14 g (Newlands some instances, mortality, in the sparsely pop-
1974a, 1978a). It also includes the world's ulated arid to semi-arid western regions of
only diurnal buthid, P. villosus (Newlands southern Africa (Muller 1993; Bergman
1974a; Hariegton 1982). 1997a, b). However, the more densely popu-
Parabuthus displays a classic “arid corri- lated mesic eastern regions are devoid ofPar-
dor” pattern of distribution (Balinsky 1962), abuthus (Prendini 1995) and the incidence of
occurring in two disjunct geographical re- scorpion envenomation in southern Africa is
gions: (1) arid southwestern Africa; (2) arid therefore lower than might be expected, given
northeastern Africa and the Arabian Peninsu- the abundance of medically important scor-
la. During past periods of increased aridity, pion species in the region.
such as the Pliocene and the Upper Pleisto- The medical importance ofParabuthus es-
cene (Tankard & Rogers 1978; Lancaster pecially P. granulatus and P transvaali,cus
,
1981, 1984; Ward et al. 1983), these two areas has received increasing attention during the
were contiguous, allowing dispersal or range past 25 years (Newlands 1974a, 1978a; New-
&
expansion of arid-adapted taxa between them. lands Martindale 1980; Petersen 1987; Hill
As evidence for their arid requirements, the 1990; Saunders & Morar 1990; Lee 1991;
southern African species of Parabuthus are Muller 1993; Bergman 1995, 1997a, 1997b;
today restricted to areas receiving less than Swells et al. 1997; DeBoet et al. 1998; Tytgat
109
110 THE JOURNAL OF ARACHNOLOGY
2 6 10
P. villosus
1 12 0 11
PRENDINI—SYSTEMATICS OF SOUTHERN AFRICAN PARABUTHUS 111
et al. 1998; Dyason et al. 2002; Huys et al. with political instability, hampers an adequate
2002). In comparison, attention to the system- revision of the Parabuthus fauna of that re-
atics of these scorpions has lagged behind. gion.
The most significant recent contribution to the No existing publication has synthesized the
taxonomy ofParabuthus was produced by La- taxonomic status ofall described southern Af-
moral (1979), who revised eleven species oc- rican Parabuthus species and subspecies, or
curring in Namibia, described three new spe- provided information on the whereabouts of
cies, and proposed eight synonyms. Probst type specimens. Two species, viz. P. calvus
(1973) reviewed the two species occurring in Purcell 1898 and P. planicauda (Pocock
Kenya and Tanzania, and Vachon (1979) and 1896), have not been revised since their orig-
Sissom (1994) reviewed the type species, P. inal description. Confusion surrounds the sta-
leiosoma (Ehrenberg 1828), in the Arabian tus ofP. brachystylus Lawrence 1928, P.flav-
Peninsula. Newlands & Martindale (1980) and idus Pocock 1899 and P. neglectus the first
,
FitzPatrick (1994) reviewed the Parabuthus two of which are currently regarded as sub-
species occurring in Zimbabwe. Kovarrk species of P. villosus and P. mossambicensis
(2003b) reviewed the Parabuthus species of (Peters 1861), respectively. The status of the
Djibouti, Eritrea, Ethiopia and Somalia, de- remaining subspecies is also uncertain.
scribed a new species and proposed seven new This contribution addresses these gaps in
synonyms. Additional contributions to Para- the knowledge of southern African Parabu-
buthus systematics in southern Africa were thus by updating the taxonomy and providing
produced by Newlands (1970), Eastwood a key to the valid species of the region. The
(1977), Lamoral (1977, 1980), Harington diagnosis ofeach species is revised, its known
(1984) and Prendini (2000, 2001a, 2003). distributional range summarized, and notes on
Parabuthus currently includes 27 species its ecology and conservation provided. Para-
and eight subspecies, four of which are nom- buthus calvus, P. capensis and P. planicauda
& &
inotypic (Lamoral Reynders 1975; Fet are redescribed and their distributions
Lowe 2000; Kovafik 2003b). Twenty of these mapped. A cladistic analysis of the southern
species and all eight subspecies are endemic African species, including five of the eight
to the southern African region. The recent dis- northeastern African and Arabian species,
covery of a new species in Namibia (Prendini based on morphological data, has been pub-
2000), and the synonymy ofP. neglectus Pur- lished elsewhere (Prendini 2001a, 2003; Fig.
cell 1899 with P. capensis (Ehrenberg 1831), 1). A larger study, involving a simultaneous
proposed here, retains the total number of analysis of morphological and molecular
southern African species at 20. (DNA sequence) data, is underway (Prendini,
This contribution focuses on the southern Esposito & Wheeler in prep.).
African species of Parabuthus for the follow-
METHODS
ing reasons. (1) Parabuthus is more diverse in
southern Africa than in northeastern Africa In the present contribution, nomenclatural
and Arabia. (2) The most venomous species emendations are based primarily upon the ex-
occur in southern Africa. (3) A limited quan- amination of type material. However, taxo-
tity ofcolonial-era material from the countries nomic decisions presented here are not de-
of greatest Parabuthus diversity in northeast- rived solely from examination of the types.
ern Africa (particularly Somalia), coupled Additional specimens were examined in all
—
Figure 1. The optimal tree obtained by an analysis (Prendini 2003) of 53 morphological characters
scored for 25 species of Parabuthus Pocock 1890 (and two outgroup taxa). This topology was retrieved
under weighting regimes that maximized fit and minimized length (length: 132 steps; fit: 59 %; Cl: 44;
RI 74). Zero-length branches are collapsed. Solid bars indicate uniquely derived apomorphic character
states, whereas empty bars indicate parallel derivations of apomorphic states under ACCTRAN optimi-
zation. The number above each bar gives the character number, whereas the number below gives the
character state. Branch support values of nodes are provided below branches. Refer to Appendix 1 for
character descriptions.
.
PRENDINI—SYSTEMATICS OF SOUTHERN AFRICAN PARABUTHUS 113
cases, in order to assess the extent of geo- diagnosable based on somatic characters, and
graphical variation and Identify characters that thus represent valid species in the diagnostic
are consistent across the distributional range. sense. A classic example is Scorpio rnaurus
Species are delimited here in accordance Linnaeus 1758 (family Scorpionidae Latreille
with the phylogenetic species concept, where 1802), with 19 currently accepted subspecies,
a species is defined as aminimum diagnosable each allopatric or parapatric, without inter-
unit (Nelson & Plateick 1981; Cracraft 1983, mediates. Neither Vachon (1952) nor Levy &
1989; Wheeler & Nixon 1990; Nixon & Amitai (1980) were able to perceive diagnos-
Wheeler 1990; Davis & Nixon 1992; De Pin- tic characters for species delimitation in S.
na 1999). As such, all subspecies are consid- maurus. Nonetheless, these authors were able
ered junior synonyms of their respective spe- to differentiate subspecies and describe them
cies unless they can be consistently separated in the absence of more “reliable” characters.
based on somatic characters. In that case, they Vachon (1952) even provided a key to the
represent valid species in the diagnostic sense subspecies, by means of which they may be
and are elevated accordingly. Note that this consistently diagnosed. Levy & Amitai (1980)
view differs from traditional usage ofthe sub- provided conclusive evidence that Scorpio
species category in scorpion systematics, in maurusfuscus (Ehrenberg 1829) and S. mau-
which the biological species concept, accord- ruspalmatus (Ehrenberg 1828) are ecological
ing to which species are defined based on re- species (seesu Van Valen 1976), each adapted
productive isolation (Dobzhansky 1937; Mayr to slightly different substrata. No evidencehas
1963), still prevails. Unlike most insects and thus far been presented to suggest that the
spiders, scorpion genitalia seldom provide morphological variation in S. maurus is clinal,
sufficient characters at the species level (al- a basic tenet of the hypothesis that this is a
though there are notable exceptions, e.g.. widespread polymorphic species. The distri-
Bothriuridae). For example, Lamoral (1979) bution of the subspecies of S. maurus con-
and FitzPatrick (1994) found no discrete di- forms to the typical distributional pattern of
agnostic differences in the hemispermato- localized endemics exhibited by other speci-
phores of ten species of Parabuthus, all of ose African scorpionid genera such as Pan-
which are diagnosable based on consistent so- dinus Thorell 1876 and Opistophthalmus C.L.
matic differences. Presumably, differences in Koch 1837.
the specific-mate recognition system (Paterson Personally collected specimens were found
1985), i.e., pre-mating isolation mechanisms by turning stones during the day, or by ultra-
(Mayr 1963), of scorpions occur at the behav- violet (UV) light detection at night (Honet-
&
ioral or chemical level (e.g., see Gaffin schlager 1965; Stahnke 1972a; Sissom et al.
Brownell 1992) rather than at the genitalic 1990). A portable UV lamp, comprising two
level. Because ofthis common paucity ofgen- mercury-vapor tubes attached to a chromium
italic variation, taxonomists have often resort- parabolic reflector and poweredby arecharge-
ed to subspecies for delimiting somatic vari- able 7 Amp/hr, 12 V battery, was used for
ation in complexes ofclosely related scorpion fieldwork. A portable Garmin® GPS II Plus
species. For example, Harington (1984:404- device was used for recording the geographi-
405) regarded P. brachystylus as a subspecies cal coordinates of collection localities in the
ofP. villosus on the grounds that morphology field.
“has a strictly secondary role as a criterion of Depositories for specimens are abbreviated
species rank”, while acknowledging that this as follows: AMGS = Albany Museum (Gra-
“is a partly arbitrary decision”. In reality, hamstown. South Africa); AMNH = Ameri-
many (probably most) scorpion subspecies are can Museum of Natural History (New York,
—
Figures 2-9. Diagnostic characters of Parabuthus species. 2-7. Metasomal segments I and II, dorsal
aspect, illustrating stridulatory surface. 2. Parabuthus mossambicensis (Peters 1861). 3. Parabuthus ca-
pensis (Ehrenberg 1831). 4. Parabuthus kraepelini Werner 1902. 5. Parabuthus stridulus Hewitt 1913. 6.
Parabuthus transvaalicus Purcell 1899. 7. Parabuthus villosus (Peters 1862). 8-9. Metasomal segment I,
lateral aspect. 8. Parabuthus villosus. 9. Parabuthus transvaalicus Scale bars = 1 mm.
114 THE JOURNAL OF ARACHNOLOGY
NY); BMNH = The Natural History Museum coverage ofsand systems was created by clip-
(London, UK); CASC = California Academy ping and merging relevant polygons extracted
GNME =
of Sciences (San Francisco, CA); from a coverage ofthe geology ofAfrica pro-
Goteborgs Naturhistoriska Museet (Goteborg, vided by the Department of Marine Geosci-
Sweden); MCZ = Museum of Comparative ence, University ofCape Town, withpolygons
Zoology, Harvard University (Cambridge, extracted from a coverage of Namibian land-
MA); MHNC = Musee d’Histoire Naturelle forms from the Namibian National Biodiver-
MMKZ
(La-Chaux-de-Fond, Switzerland); -• sity Task Force (Barnard 1998), downloaded
McGregor Museum (Kimberley, South Afri- from their website: http://www.dea.met.gov.
ca); MNHN — Museum National d’Histoire na/programmes/biodiversity/countrystudy.htm.
Naturelle (Paris, France); NHMB = Naturhis- In order to create a point locality geograph-
torisches Museum Basel (Basel, Switzerland); ical dataset for mapping distributional ranges,
NHMW
- Naturhistorisches Museum Wien all records ofsufficient accuracy wereisolated
(Vienna, Austria); NHRS = Naturhistoriska from the material examined. Only a small pro-
NMNW =
Riksmuseet (Stockholm, Sweden); portion of the records were accompanied by
National Museum ofNamibia (Windhoek, Na- geographical coordinates or quarter-degree
mibia); NMSA = Natal Museum (Pietermar- squares (QDS), usually entered by the collec-
itzburg, South Africa); SAMC = South Afri- tor or subsequently added by the curator or
can Museum (Cape Town, South Africa); collections manager. These were checked for
TMSA = Transvaal Museum (Pretoria, South accuracy and an attempt was made to trace
Africa); ZMHB = Zoologisches Museum, coordinates for as many of the remaining re-
Universitat Humboldt (Berlin, Germany). cords as possible, by reference to gazetteers,
Illustrations were produced using a Leica the official 1:250 000 and 1:500 000 topo-ca-
MZ16 stereomicroscope and camera lucida. dastral maps of South Africa and neighboring
Photographs were taken using a digital pho- countries published by the Government Print-
tomicrography system (Microptics® ML- er, and the GEOnet Names Server: http://
1000). Measurements were made with Mitu- 164.214.2.59/gns/html/cntry_files.html.
toyo® digital calipers (model NTD12-6"C). Names of Namibian regions and magisterial
Color designation follows Smithe (1974, districts and South African provinces and
1975, 1981), trichobothrial notation follows magisterial districts listed in the material ex-
Vachon (1974), sternum terminology follows amined follow the most recent system (post-
&
Soleglad Fet (2003) and measurements fol- 1994).
low Stahnke (1970), Eastwood (1977) and La-
Genus Parabuthus Pocock 1890
moral (1979). Morphological terminology fol-
lows Couzijn (1976) for the segmentation of Buthus (Parabuthus) Pocock 1890: 124, 125; type
legs, Hjelle (1990) and Sissom (1990) for the species by original designationAndroctonus leio-
segmentation of pedipalps, Prendini (2000, soma Ehrenberg 1828 [= Parabuthus leiosoma
2001a, 2003) for metasomal carinae, and (Ehrenberg 1828)].
Stahnke (1970), Lamoral (1979), Sissom Heterobuthus Kraepelin 1891: 63-68; type species
by subsequent designation (Kraepelin 1895: 78),
(1990) and Prendini (2001a) for remaining
Androctonus leiosoma Ehrenberg 1828 [= Par-
characters.
abuthus leiosoma (Ehrenberg 1828)] (synony-
Distribution maps forP. calvus, P. capensis mized by Thorell 1893: 366).
and P. planicauda were produced using Parabuthus Kraepelin 1895: 82; Pocock 1895:
:
ArcView GIS Version 3.2 (Environmental 309; Laurie 1896: 131; Lonnberg 1897; 194;
Systems Research Institute [ESRI], Redlands, Kraepelin 1899: 28; Birala 1917a: 164; Birola
CA), by superimposing point locality records 1917b: 55; Pavlovsky 1924: 77; Werner 1934:
on coverages depicting the topography (500 269; Kastner 1941: 232; Bucherl 1964: 57; Stahn-
m contour interval), major sand systems and ke 1972b: 130; Lamoral 1979: 560, 561; Fr&ancke
ppoolgirtaicpahlicbocuonndtaoruirescoovfersaoguethweransAcfrreiactae.dAfrtoom- 11998952:; K1o1,va1n6;kS1i9s98s:om11169;901:17;10F2e;tN&enLioliwne200F0e:t
the GTOPO30 raster grid coverage, obtained 200; Prendini 2001a: 13-35; Dyason et al. 2002:
768-773; Prendini 2003: 5, 19-22.
from the website ofthe U.S. GovernmentPub- —
lic Information Exchange Resource: http:// Diagnosis. The genus Parabuthus is most
edcdaac.usgs.gov/gtopo30/gtopo30.htmL A closely related to the Afrotropical buthid gen-
)
PRENDINI—SYSTEMATICS OF SOUTHERN AFRICAN PARABUTHUS 115
—
era Grosphus Simon 1888 and JJroplectes Pe- Distribution. Angola, Botswana, Djibouti,
ters 1861 (Prendini 2001a, 2004). All species Egypt, Ethiopia, Kenya, Mozambique, Namib-
of Parabuthus with one exception (viz. P. ia, Saudi Arabia, Somalia, South Africa, Su-
,
distridor Lamoral 1980), are readily distin- dan, Tanzania, Yemen, Zambia, Zimbabwe.
guished from Grosphus Uroplectes and other Twenty-eight species are currently recognized,
,
buthid genera by the presence of a distinct 20 of which are endemic to the southern Af-
stridulatory surface, composed of fine to rican region, whereas eight are endemic to
coarse granules, sometimes forming trans- northeastern—Africa and the Arabian Peninsula.
verse ridges, on the dorsal surfaces of tergite Ecology. All species of Parabuthus are
VII, and metasomal segments I, II and, to a adapted for a burrowing existence (Newlands
lesserextent, III (Newlands 1974a, 1978a; La- 1974b, 1978a, b). The thickened metasoma,
moral 1977, 1979, 1980; Figs. 2-7). This especially the processes on the ventrolateral
stridulatory surface is used to make an audible carinae of metasomal segment V, is used to
sound (stridulation) when the scorpion is loosen the soil, whereas the anterior two pairs
alarmed by repeatedly scraping the aculeus of legs are used to scrape and rake the loos-
across the granules orridges (Dumortier 1964; ened soil out of the burrow (Eastwood 1977,
Newlands, 1974a, 1978a; McCormick & Polis 1978b; Lamoral 1979; Prendini 2001b). Bur-
1990). Additionally, all species ofParabuthus rows are constructed in open ground, at the
(including P. distridor) are characterized by base of shrubs or under stones (Newlands
the following unique combination of charac- 1974b, 1978a, b; Eastwood 1977, 1978b; La-
ters: carapace without granular carinae; me- moral 1979; Newlands & Martindale 1980;
sosomal tergites with only a single, weakly Harington 1984; Prendini 2000, 2001a,
developed median carina; proximal median la-
2001b).
mella ofpectines, enlarged and lobate (dilate) Many Parabuthus species are morphologi-
in female (Fig. 11; P. granulatus and P. ka- cally specialized for life on particular substra-
laharicus Lamoral 1977 are exceptions in this ta. Some (e.g., P. distridor P. kalaharicus and
regard); basitarsi of legs I and II with macro- P. kuanyamarum Monard ,1937) are restricted
setal combs (Fig. 12); metasomal segment IV, to the sand dune systems of the Namib and
ventrosubmedian carinae becoming obsolete
Kalahari deserts, and exhibit ecomorphologi-
distally (Figs. 14, 15); metasomal segment V,
cal adaptations to facilitate locomotion and
ventrolateral carinae comprising distinct spi-
burrowing in the soft sand (Lamoral 1977,
nose or lobate processes subdistally (Figs. 22,
1979, 1980; Prendini 2001b). Others (e.g., P.
29, 31). —
Remarks. Prior to the present contribu- planicauda and P. villosus) appear to be mor-
tion, all but two of the southern African spe- phologically specialized for inhabiting rocky
cies had been revised to some extent. In con- areas (Prendini 2001b).
trast, only one ofthe northeastern African and Adaptive radiation in Parabuthus may be re-
Arabian species, P. leiosoma has been par- lated to the evolution ofburrowing behavior in
tially revised (Sissom 1994). The limited the genus. According to the “EffectHypothesis
quantity of material available for most north- of macroevolution” (Vrba 1980), repeated al-
eastern African species (except P. leiosoma lopatric speciation in Parabuthus is the pre-
remains an obstacle for revising the species in dicted outcome of vicariance, promoted by
that region, given the extent of geographical stenotopic habitat requirements, in this case
variation in Parabuthus species. “substratum specialization” (Prendini 2001b).
ILLUSTRATED KEY TO IDENTIFICATION OF THE SOUTHERN AFRICAN SPECIES OF
PARABUTHUS POCOCK 1890
1. Metasomal segment IV, ventrosubmedian and ventrolateral carinae absent (Fig. 13) .... . 2
Metasomal segment IV, ventrosubmedian and ventrolateral carinae present, but ventrosub-
median carinae becoming obsolete distally (Figs. 14, 15) 7
2. Pedipalp chela asetose; trichobothrium dt in line with or distal to et (Figs. 55, 56) .....
P. calvus
Pedipalp chela covered in setae; trichobothrium dt proximal to et (Figs. 42-45) 3
116 THE JOURNAL OF ARACHNOLOGY
3. Metasomal segments I and II, dorsal stridulatory surface absent, ventrosubmedian and
ventrolateral carinae absent (Fig. 28) P. distridor
Metasomal segments I and II, dorsal stridulatory surface present (Figs. 2-7), ventrosub-
median and ventrolateral carinae present (Figs. 25—27) 4
4. Metasomal segments II and III, distal section ofventrolateral carinae composed ofstrongly
elevated, crescent-shaped tubercles, forming abroad U-shaped pattern (Fig. 27); metasomal
segment V, ventrolateral carinae with lobate processes (broad and presenting a flat surface
apically) subdistally (Fig. 29) P. brevimanus
Metasomal segments II and III, distal section of ventrolateral carinae costate or composed
of isolated, round granules, not forming a U-shaped pattern (Figs. 25, 26); metasomal
segment V, ventrolateral carinae with spinose processes (narrow, conical or flattened, and
tapering apically) subdistally (Fig. 30) 5
5. Pedipalp chela smooth and shiny (Fig. 46); carapace (female) with smooth areas
P. kuanyamarum
Pedipalp chela granular (Figs. 43-45); carapace (female) entirely granular ............ 6
6. Pedipalp chela movable finger, short, compared with manus (measured along ventroexternal
carina), length finger/length carina: ± 1.50 (Fig. 43); chela manus (adult male) incrassate
(bulbous or swollen) (Fig. 43) P. gracilis
Pedipalp chela movable finger, long, compared with manus (measured along ventroexternal
carina), length finger/length carina: 1.70-2.00 (Figs. 44, 45); chela manus (adult male)
slender (Fig. 44) P. nanus
7. Metasomal segment IV, median lateral carina absent or more weakly developed than dor-
solateral and ventrolateral carinae (Figs. 16, 17) 8
Metasomal segment IV, median lateral carina as strongly developed as dorsolateral and
ventrolateral carinae (Fig. 18) 12
8. Metasomal segment II, dorsal stridulatory surface reaching posterior margin (Figs. 2-6);
width of telson vesicle equal to distal width of metasoma V (Figs. 29, 30, 32) 9
Metasomal segment II, dorsal stridulatory surface not reaching posterior margin (Fig. 7);
telson vesicle narrower than distal width of metasoma V (Fig. 31) 11
9. Pedipalp chela manus covered in setae; metasomal segment V, dorsosubmedian carinae
present but weakly developed (Figs. 20, 23) P. namibensis
Pedipalp chela manus asetose; metasomal segment V, dorsosubmedian carinae absent (Figs.
19, 22) 10
10. Metasomal segment II, dorsal stridulatory surface composed of granules (Figs. 3, 4, 6, 7);
telson vesicle dorsoproximal surface very shallowly excavated along longitudinal half(Fig.
34) P. laevifrons
Metasomal segment II, dorsal stridulatory surface composed of granular ridges (Fig. 5);
telson vesicle dorsoproximal surface deeply excavated along longitudinal half (Fig. 33)
P. stridulus
11 Pedipalp chela, trichobothrium dt in line with or distal to et (Figs. 39, 40); metasomal
.
segment IV, ventrosubmedian carinae becoming obsolete in distal third of segment (Fig.
15) P. granulatus
Pedipalp chela, trichobothrium dt proximal to et (Figs. 37, 38); metasomal segment IV,
ventrosubmedian carinae becoming obsolete in proximal third of segment (Fig. 14)
P. kalaharicus
12. Metasomal segment II, posterodorsal edge extended forwards in V-shape (Fig. 3) ..... 13
Metasomal segment II, posterodorsal edge straight (Figs. 2, 4-7) .................. 14
13. Pedipalp chela fingers fit together with distinct proximal gap, when closed (Fig. 41); telson
aculeus short, sharply curved (Fig. 36); chela manus (adult male) slender (Figs. 44, 55,
83) P. muelleri
Pedipalp chela fingers fit together without gap, when closed (Figs. 69, 70); telson aculeus
long, gently curved (Fig. 35); chela manus (adult male) incrassate (Fig. 69) .... P. capensis
14 Metasomal segment V, ventrolateral carinae with lobate processes (Figs. 22, 29, 31) ... 15
.
Metasomal segment V, ventrolateral carinae with spinose processes (Figs. 24, 30, 32) 17
. .
PRENDINI—SYSTEMATICS OF SOUTHERN AFRICAN PARABUTHUS 117
15. Metasomal segment II, dorsal stridulatory surface composed of ridges (Fig. 2)
P. mossamhicensis
Metasomal segment II, dorsal stridulatory surface composed of granules (Figs. 3, 4, 6, 7)
16
16. Metasomal segments I and II, dorsal stridulatory surface extended anteriorly (Fig. 8); chela
manus (adult male) slender (Fig. 83) P. planicauda
.
Metasomal segments I and II, dorsal stridulatory surface truncated anteriorly (Fig. 9); chela
manus (adult male) incrassate (Figs. 37, 39, 43, 46, 69) ................... P. kraepelini
Metasomal segment V, ventrolateral carinae converging distally (Fig. 30); coloration pale,
although metasomal segments IV and V and telson may be infuscated .......... P. raudus
Metasomal segment V, ventrolateral carinae subparallel to diverging distally (Figs. 29, 31,
32); coloration dark (pedipalps and legs may be pale) 18
17. Metasomal segments I and II, dorsal stridulatory surface extended anteriorly (Fig. 8); me-
tasoma densely setose; metasoma I wider than IV P. villosus
Metasomal segments I and II, dorsal stridulatory surface truncated anteriorly (Fig. 9);
metasoma moderately setose; metasoma I narrower than IV ...................... . 19
18. Metasomal segment II, dorsal stridulatory surface reaching posterior margin (Fig. 6); col-
oration of pedipalps and legs dark, not contrasting markedly with dark pro-, meso- and
metasoma P. transvaalicus
.
Metasomal segmentII, dorsal stridulatory surface notreaching posteriormargin ofsegment
(Fig. 7); coloration ofpedipalps and legs pale, contrasting markedly with dark pro-, meso-
and metasoma P. schlechteri
Parahuthus brevimanus (Thorell 1876) separated from all other species ofParahuthus
Figs. 27, 29 by means of the following combination of
Buthus brevimanus Thorell 1876b: 110-113. characters: small adult size, carapace length
Heterobuthus brevimanus Kraepelin 1891: 69. 2.5-5.0 mm; pedipalp chela with trichoboth-
:
Parahuthus brevimanus: Pocock 1895: 309; Krae- rium dt situated proximal to et; metasomal
pelin 1899: 32; Purcell 1901: 149-151; Kraepelin segments slender (length IV/width IV: 1.7—
1908: 250; Hewitt 1913: 146; Kraepelin 1914: 2.11); metasomal segment IV with median lat-
111, 112; Hewitt 1918: 104, 176; Fage 1925: eral, ventrolateral and ventrosubmedian cari-
189; Lawrence 1927: 72; Lawrence 1928: 269; nae absent; metasomal segment V with
Werner 1936: 177; Monard 1937: 256, 257;
dorsosubmedian, dorsolateral and ventrome-
Lawrence 1955: 225; Lawrence 1959: 383
Lawrence 1962: 220; Lamoral & Reynders 1975:; dian carinae absent.
514, 515; Lamoral 1979: 561-566, figs. 87-94, Parahuthus brevimanus is most easily con-
97-99 Kovaffk 1998: 116; Fet & Lowe 2000: fused with P. gracilis, with which it is sym-
;
201; Prendini 2001a: 17; Prendini 2001b: 137; patric in the northwestern part of its distribu-
Dyason et al. 2002: 769; Prendini 2003: 20. tional range (Erongo and Kunene regions,
Parahuthus cristatus Pocock 1901: 284, 285 (syn- Namibia). However, P. brevimanus may be
onymized by Kraepelin 1908: 250; Lamoral separated from P. gracilis, as well as P. dis-
1979: 561); Pocock 1902: 367, 368. tridor, P. kuanyamarum and P. nanus, on the
—
Type material. Buthus brevimanus: Lec- basis of the following combination of char-
totype 9 (GNME) [designated by Lamoral acters: surface of median ocular tubercle
(1979)], “S. Afrika”, 28.xi.1864 [not (male, female) and surrounding surfaces of
23.xi.1864], C.J. Andersson. Paralectotype S carapace (female), smooth and shiny; pedipalp
(NHRS) [designated by Lamoral (1979)], chela manus smooth; metasomal segments I—
“Caffraria”, 1840-1845, J. Wahlberg. IV with dorsosubmedian carinae present, but
Parahuthus cristatus: Holotype 9 obsolete; metasomal segments II and III, distal
(BMNH), “Co—ngo” [probably Angola]. section of ventrolateral carinae, and poster-
Diagnosis. Parahuthus brevimanus is oventral margin composed of strongly elevat-
basal to the clade comprising the P. distridor- ed, crescent-shaped tubercles, forming abroad
P. kuanyamarum and P. gracilis-P. nanus sis- U-shaped pattern; metasomal segment IV, an-
ter groups (Fig. 1). The five species may be teroventral margin demarcated by a transverse
—
118 THE JOURNAL OF ARACHNOLOGY
—
Figures 10-12. Diagnostic characters ofParabuthus species [modified from Lamoral (1979)]. 10, 11.
Dextral pecten of 9, ventral aspect. 10. Uroplectes carinatus (Pocock 1890). 11. Parabuthus gracilis
Lamoral 1979. 12. Parabuthus stridulus Hewitt 1913, distal segments of dextral leg I, dorsal aspect,
illustrating macrosetal combs along retrolateral edge. Scale bars = 1 mm.
row of 5-6 strongly elevated crescent-shaped P. cristatus and erroneously claimed that P.
tubercles; metasomal segment V with distally cristatus wa,s a —new synonym.
diverging ventrolateral carinae, each compris- Distribution. Recorded from Angola, Na-
ing several prominent lobate processes sub- mibia, and South Africa. The distribution of
distally. — this species extends across the Orange River.
Remarks. Purcell (1901) listed P. crista- Ecology. Parabuthus brevimanus is a
tus as ajunior synonym ofP. brevimanus, but semi-psammophilous species, inhabiting
placed a question mark next to the synonymy, semi-consolidated to consolidated sandy and
as he had not examined the type specimens. gritty substrata, where it excavates burrows in
Kraepelin (1908) confirmed the synonymy by open ground and at the base of shrubs and
comparing the type specimens. Most subse- grass tufts. Comb-like rows of long macrose-
quent authors adopted the synonymy (Krae- tae (“sand combs”) on the retrolateral mar-
pelin 1914; Hewitt 1918; Lawrence 1955), but gins of the basitarsi of legs I and II indicate
Lamoral & Reynders (1975) continued to re- a semi-psammophilous ecomorphotype. Me-
gard P. cristatus as a distinct species. Lamoral tasomal segments IV and V are mostly lacking
(1979) later revised P. brevimanus re-exam- carinae.
,
ined the type specimens ofP. brevimanus and Parabuthus brevimanus is syntopic with P.
